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A plumber's-eye view of xylem water transport in

woody plants
a

Jordi Martnez-Vilalta & Josep Piol

CREAF/Ecology Unit , Universitat Autnoma de Barcelona , Barcelona, Spain

Published online: 13 Dec 2010.

To cite this article: Jordi Martnez-Vilalta & Josep Piol (2004) A plumber's-eye view of xylem water transport in woody
plants, Journal of Biological Education, 38:3, 137-141, DOI: 10.1080/00219266.2004.9655923
To link to this article: http://dx.doi.org/10.1080/00219266.2004.9655923

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Practical
A plumber's-eye view of
xylem water transport in
woody plants

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Jordi Martfnez-Vilalta and Josep Pihol

CREAF/Ecology Unit, Universitat Autonoma de Barcelona, Barcelona, Spain
We present a practical for university-level students aimed at measuring and comparing xylem hydraulic
properties of co-existing plant species. After sampling branches of several woody species in the field, their
main hydraulic properties were measured using a simple set-up. Hydraulic conductivity (Kh) was calculated as
the ratio between water flow through a plant segment and the pressure gradient driving the flow. The percent
reduction in conductivity due to xylem embolism (i.e. air-filled conduits) was estimated by comparing Kh before
and after flushing the measure segments to remove all native embolism. Raw hydraulic conductivity was
standardised by cross-sectional wood area or supported leaf area to obtain more meaningful measures of
conducting capacity. The results showed differences among study species, particularly between conifers and
angiosperms. These differences are briefly discussed in terms of wood anatomy and the general biology of
the species. Overall the practical provides a good opportunity for students to appreciate the main aspects of
xylem water transport and the constraints it imposes on plant water relations.
Key words: Ecophysiology, Hydraulic architecture, Hydraulic conductivity, Plant water transport, Xylem
embolism.

Introduction
Plants transport enormous amounts of water from the soil to the
atmosphere. Of the water absorbed by roots only about 5% is
used in metabolism and growth, whereas the remaining 95% is
lost through leaves in transpiration. As an example, a single isolated
tree may transpire 200-400 litres of water per day (Kozlowski
and Pallardy, 1997). Transpiration is intimately linked to carbon
acquisition, since both the flow of water out of the leaf and the
flow of carbon dioxide into the leaf occur through the same
pores (i.e. stomata). At the same time, the huge quantity of
water transported from the soil to the leaves is likely to assist the
nutrient supply of above ground tissues. How can plants pull up
these amounts of water to heights up to more than 100 metres?
According to the nearly-hegemonic cohesion-tension theory,
solar radiation provides the energy source for water evaporation
in leaves, which together with the cohesive strength of liquid
water makes the transport along the gradient of water potential
possible (Steudle, 1995).
Inside plants water is transported mainly through the xylem,
a tissue containing a network of conduits (walls of dead cells)
interconnected through porous membranes. A corollary of the
cohesion-tension theory is that water transport in the xylem
should take place at pressures substantially lower than atmos
pheric (i.e. under tension). However, at ambient temperature
(around 20C) liquid water at pressures below +2.3kPa is
thermodynamically unstable. As xylem pressures become
increasingly negative, a point is eventually reached at which
water evaporates (cavitation) and the affected conduit is filled
with air from the surrounding tissue (embolism). Inter-conduit
Journal of Biological Education (2004) 38(3)

pit membranes act as filters preventing the propagation of

embolisms. However, if xylem pressure is low enough to suck air
through inter-conduit pores (the required tension depends on
pore size) generalised embolism occurs (Zimmermann, 1983).
Embolised conduits no longer contribute to water transport
unless they are refilled with liquid water.
High levels of xylem embolism reduce the capacity of plants
to transport water and when maintained over long periods of time
can cause branch or plant dieback. Therefore, generalised embolism
is something to be avoided by plants. But what environmental
conditions favour the occurrence of xylem embolism? Drought
is clearly one of them, because the lower the water content of
the soil the lower the water potential needed by plants to extract
and transport water and, thus, the higher the amount of embolism
in the xylem. As a result, vulnerability to xylem embolism is one
of the key components of drought resistance in plants (Hacke
and Sperry, 2001). Freezing can also induce embolism: since the
solubility of air is lower in ice than in liquid water, freezing
forces air out of the xylem sap solution, forming bubbles. On
thawing, these bubbles can either re-dissolve or nucleate cavitation,
depending on the size of the conduit (larger conduits being
more vulnerable). There is evidence supporting the idea that
freezing-induced embolism limits the distribution in cold habitats
of many plant species (Hacke and Sperry, 2001).
In this paper we present a practical for university students
aimed at measuring and comparing the hydraulic properties of
several co-existing plant species. Previous studies have stressed
the faulty understanding of transpiration or plant water transport
of secondary (Barker, 1998) or university-level students
137

Xylem water transport

Martinez-Vilalta and Pinol

(Martinez-Vilalta et al). Although it is clear that a

minimum knowledge of the mechanism of sap ascent is required
to fully understand the experiments reported in this paper, in our
opinion they provide a good opportunity for students to deepen
dieir knowledge of die cohesion-tension mechanism and appreciate
some of its basic implications. Despite the fact that the study of
the hydraulic properties of plant xylem as described in this
paper is relatively new (it was systematised during the 1980s,
when some of the methods we present here were developed) it
is already recognised as one of the key aspects of plant water
relations in major textbooks on plant ecology and ecophysiology
(five pages out of 23 are devoted to this subject in the chapter
on plant water relations in Crawley [1997], and 11 out of 50 in
Lambersetal [1998]).

Consequently, the length of sampled branches should be enough

that none (or very few) of the conduits embolised during
collection extend to measured segments. Although 40-50 cm is
normally enough, some species (e.g. typically Quercus spp.) have
very long vessels and require the sampling of longer branches. It is
possible to know if there is any single vessel extending through
out a given wood segment by injecting pressurised air (at around
60 kPa) into one of the ends of the segment and submerging the
other end in water: bubbles will appear at the
submerged end only if there is at least one individual vessel
crossing the entire segment (Ewers and Fisher, 1989). Immediately
after cutting, branches were sealed in plastic bags and carried to
the laboratory. Travel time was about four hours. Once in the
laboratory samples were conserved at 2C until they were
measured within 12 hours.

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Methodology
All sampling and measurements were carried out by 12 groups
(20-25 people each) of second year students of biology at the
Universitat Autonoma de Barcelona (Spain) as a part of a one-week
course on practical ecology. The particular practical we present
here lasted for one-two days, depending on the duration of the
subsequent class discussion.
Field

sampling

The first step consists of selecting the group of woody species

that will be studied. In the particular example we report here all
species (see Table 1) were sampled during early autumn in a
Quercus ilex forest from NE Spain (at La Castanya Biological
Station [Montseny], 41 46' N, 2 21' E). All individuals (N =
three - 11 individuals per species) were sampled in the same
valley, widiin less than 500 metres. The particular species sampled
will obviously depend on the region where the practical is
conducted, but in principle any community with more than
three or four different woody species is suitable. It is advisable
to select species with contrasting life strategies, for example by
first providing students with information on the general biology
of the dominant woody species in the area and then letting
them decide the target species in terms of previous information
and personal preferences.
Fully exposed, terminal branches at least 1 m long were
collected from four-five different individuals per species. The
sampled branch that will be used in all hydraulic measurements
should contain a central segment about 20 cm long and
0.5-1 cm in diameter, with no or very few side branches.
Because xylem pressures are lower than atmospheric, cutting
introduces air (embolism) into severed xylem conduits.

Hydraulic

properties

Hydraulic conductivity (KJ is defined as the ratio between a

water flow (7) and the pressure gradient required to produce it
(Vp), and is, thus, a measure of capacity to transport water. We
measured xylem hydraulic conductivity following Sperry et al
(1988). Segments about 20cm long and with a diameter of
0.5-1 cm were re-cut underwater from the sampled branches.
Their proximal ends were connected and clamped to a tubing
system filled with filtered (4> = 0.22pm), deionised water, and
connected to a water reservoir (see Figure 1). This reservoir was
around 60cm above the level of the branches, equivalent to a

water reservoir

tubing

height'pressure
difference

Figure 1 Diagram of the set-up we used to measure hydraulic conductivity

of branch segments.

Table 1 Main ecological and wood anatomical attributes of the study species1.
Plant species

Family

Life form

Wood type

Leaf habit

Macroclimate

Abies sp.2
Alnus glutinosa
Castanea sativa
Celtis australis
Cistus salviifolius
Hedera helix
Quercus ilex

Pinaceae
Betulaceae
Fagaceae
Ulmaceae
Cistaceae
Araliaceae
Fagaceae

Tree
Tree
Tree
Tree

Conifer
Semi-ring-porous
Ring-porous
Ring-porous
Diffuse porous
Semi-ring-porous
Diffuse porous

Evergreen
Deciduous
Deciduous
Deciduous
Summer dec.
Evergreen
Evergreen

Temperate
Temperate
Temperate
Mediterranean
Mediterranean
Temperate
Mediterranean

Shrub
Liana
Tree

'Data compiled from Baas and Schweingruber (1987).

Sampled trees correspond to an exotic Abies species (probably a hybrid between A. alba and A. pinsapo) planted in the area.

138

Journal of Biological Education (2004) 38(3)

) Xylem water transport

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pressure difference of approximately 6kPa. The resulting water

flow was measured gravimetrically, by weighing previously zeroed,
cotton-filled vials that had been attached to the distal ends of the
segments for a known period of time. Hydraulic conductivity
(Ky in m 4 MPa"1 s"1) was calculated as the ratio between the
flow through the segment and the pressure gradient (pressure
difference divided by the length of the segment]. This initial
hydraulic conductivity [Kh.) corresponds to the conditions under
which the sampled individuals were operating in the field and,
therefore, includes any native embolism they were experiencing.
In order to obtain their maximum hydraulic conductivity
{KhmaJ the segments were flushed at high pressure (600 kPa]
with water for around 60 min to remove all native embolisms.
In our case this was achieved by lifting the water reservoir to the
ceiling (about six metres above]. After flushing, the water reservoir
was lowered again and hydraulic conductivity was re-measured
exactly as before. The percentage loss of hydraulic conductivity
(PLC) due to the amount of embolism originally present in the
segments can be evaluated by comparing their hydraulic
conductivities before and after flushing:

PIC = 100- l j ^ !
\

h, max

Martinez-Vilaltaand Pinol

and between 8.91 x 10"9 m 2 MPa"1 s"1 [Abies sp.] and 1.03 x 10"6
[Castanea sativd) [KA. In both cases hydraulic conductivity was
significantly lower in Abies sp. than in any other species (see
Figures 2a and 2b]. High KL of C. sativa (Figure 2b] is related
with larger AS:AL in this species (Figure 2d), which was starting
to lose leaves at the time of measurement. There was a wide
range of inter-specific variation in estimated loss of conductivi
ty due to xylem embolism (PLC), from around 0% in Abies sp.
to > 80% in Celtis australis. Loss of conductivity was signifi
cantly lower in Abies sp. than in any other species, and larger in
C australis and Q. ilex than in A. glutinosa; the rest of species
showing intermediate values (see Figure 2c).

Equation 1
/

In the few cases in which Khi>Kh

(probably because measured
flows approached the limits of the resolution of our 0.001-g
precision balance] it was assumed that PLC ~ 0. Since hydraulic
conductivity is clearly size-dependent it is always advisable to
relate it to other branch properties, like cross-sectional wood
area or supported leaf area. Specific hydraulic conductivity
[K$, in m 2 MPa"1 s"1) was calculated as the ratio between Khmax
and cross-sectional area of the segment (without bark]; and leafspecific conductivity [Ku in m 2 MPa"1 s"1], as the quotient
between Kh max and supported leaf area. An additional variable
of interest to characterise hydraulic architecture is the ratio
between cross-sectional area and leaf area
[A^AJ
(Zimmerman, 1983]. In our case, leaf area was measured with a
LI-COR 3100 leaf area meter (LI-COR). If similar equipment is
not available it is possible to estimate mass per unit leaf area
[LMA, in mg cm 2 ] in a reduced amount of leaves and use this
value to convert total leaf weight into total leaf area.
Data

analysis

Hydraulic properties were compared among species by means

of one-way AN OVA (plus Tuckey HSD a posteriori test]. Since
none of the variables studied was normally distributed they
were normalised (logarithmic transformation for Ks, KL and
AS:AL; and arcsine transformation for PLC] prior to analysis.
Consequently the averages and confidence intervals (CI] provided
in the text and figures correspond to those calculated for the
transformed variables back-transformed to the original scale.

Results
Typical water flows through measured segments were 0.1-1 mg
s"1 for angiosperms and 0.01-0.1 mg s"1 for Abies sp. (globally 10
- 1000 mg deposited in a 15-minute measurement; enough to
be measured with a 0.001 g-precision balance]. When standard
ised according to cross-sectional area of wood and leaf area,
hydraulic conductivity ranged, respectively, between 1.44 x 10"
5
{Abies sp.] and 9.99 x 10"4 m 2 MPa"1 s"1 [Alnus glutinoso) [Ks),
Journal of Biological Education (2004) 38(3)

Figure 2 Hydraulic properties of branches of the study species, including:

specific hydraulic conductivity (K^ (a), leaf-specific hydraulic conductivity
(KJ (b), percent loss of conductivity due to xylem embolism under field
conditions (c), and wood-to-leaves area ratio (d). Column heights and error
bars correspond, respectively, to averages and 95% confidence intervals
calculated for normalised variables back-transformed to the original scale
(see 'Data analysis' section). Different letters indicate differences between
species at the 0.05 significance level. See Table 1 for species' full names.
139

) Xylem water transport

Martinez-Vilalta and Pifiol

Discussion
This practical offers several opportunities for discussion.
Measured water flows illustrate very clearly one essential property
of wood: its capacity to transport water longitudinally. Xylem
occupies the main part of the cross sectional area of a typical
stem or root and has a huge hydraulic conductivity compared to
other tissues (e.g. Ks of parenchyma is = 10~9 m 2 MPa"1 s"1
[Raven, 1993], four to six orders of magnitude lower than those
measured in this study for wood segments). Without this high
wood hydraulic conductivity terrestrial plants would never have
reached the heights we currently observe in most forests. Measured
hydraulic conductivities can be scaled up to the tree level to
provide a rough idea of the amount of water that can be trans
ported by a whole individual of the study species. If it is assumed
that measured Ks is representative of above-ground parts, the
flow of water (7) through the trunk can be obtained from:

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J = Ks A s VM*

Equation 2

If we take, for example, a 20 cm-diameter Q. ilex tree and consider

a reasonable gradient of water potential (Vi); = 0.1 MPa m"1], we
obtain that 7 = 2 . 4 x 10~6 m 3 s_l = 8.5 1 h"1. This value is reason
ably similar to the actual flows estimated for Q. ilex trees of the
same size in a similar forest (Martinez-Vilalta et al, unpublished)
and gives us an idea of the magnitude of the flow that can be
supported by the xylem.
What determines the hydraulic conductivity of the xylem? As
a first approximation xylem conduits can be assimilated to
capillaries and, consequently, it is possible to describe the water
flow across them using the Hagen-Poiseuille equation:
N

y
*

n
v
11 K,

Equation 3

i 8p

where N is die number of functional conduits, R. is die radius of die

i conduit (in m), and u, is the viscosity of water (1.002 x 10"9
MPa s at 20C). In angiosperms xylem water transport occurs
basically through relatively wide and long multicellular conduits,
termed vessels (diameter = 10-200 |xm). In conifers, which do
not have vessels, water moves dirough smaller conduits (tracheids],
typically less than 10 u,m in diameter (see Figure 3). The
dependence of hydraulic conductivity on conduit radius raised
to the fourth power (Equation 3) explains why Ks and KL were
lowest in Abies sp. (see Figure 2]. Although we have not tried it in
the practicals, it is easy to cut wood sections from the measured
segments using a hand microtome or a razor blade and, with the

aid of a microscope, obtain representative values of conduit size

and density for each study species (see Ewers and Fisher, 1989
for detailed methods]. These values can be used to estimate a
theoretical Kh max. Alternatively, representative values (if they
are known] can be provided by the tutor. Theoretical calculations
normally overestimate measured conductivities because of the
existence of resistances not accounted for in Equation 3 (e.g.
inter-conduit membranes].
The fact that initial hydraulic conductivity increased after
flushing the segments (i.e. PLC > 0) is a clear illustration of one
of the main drawbacks of the mechanism of sap ascent in plants.
In most plants xylem hydraulic conductivity under field conditions
is substantially lower than Ki
because of air-filled, non
functional conduits. What explains the presence of high levels of
embolism in some of the study species (Figure 2c]? In our case
it is probably a combination of residual summer embolism (Q.
ilex, for example is known to be highly vulnerable to droughtinduced xylem embolism; Martinez-Vilalta et al, 2002] and
freezing-induced xylem embolism, particularly in winter
deciduous species (see Table 1].
Embolism and the timing of leaf fall are known to be related:
high levels of embolism during the autumn are associated with
early leaf fall and vice versa (Hacke and Sperry, 2001]. The
apparent exception of A. glutinosa, a deciduous species with
very low PLC, arises from the fact that this species was sampled
at the bottom of the valley while all the other species were sam
pled in the mid slopes of the same valley. In the area where A.
glutinosa was sampled, soil water potentials were probably close
to 0 and, therefore, no (or little] embolism could be formed.
Words of caution
Before concluding, several words of caution are needed. It has
been shown that a considerable insight can be gained from the
consideration of plant xylem as a tissue composed of dead cells,
acting as passive conduits. However, recent studies suggest that
living parenchyma cells within the xylem may have a more
important role in water transport than previously thought, for
example in the refilling of previously embolised conduits
(Holbrook and Zwieniecki, 1999].
Moreover, there is some degree of controversy on the validity
of the cohesion-tension theory (Zimmermann et al, 2000],
although the vast majority of researchers agree on its main
aspects. Finally, in this paper we have adopted a relatively sim
ple view, focusing on xylem water transport, without explicitly
considering other aspects, notably soil-root and leaf-atmosphere
interfaces, which together with xylem transport determine
whole plant water economy.

Acknowledgements
We would like to thank the students of second year of biology
at the Universitat Autonoma de Barcelona (academic years
2001 - 2002 and 2002 - 2003], who obtained all the results we
present here and improved the practical with their suggestions.
The comments by two anonymous reviewers substantially
improved the manuscript.
Figure 3 Photographs of representative cross-sections of the xylem of an
angiosperm (left, Quercus ilexj and a conifer (right, Pinus pineaj. Both
photographs are at the same scale (white bar length = 100 fim). Note the
difference in the general structure of the xylem and, particularly, the much
wider conduits in the angiosperm.
140

Journal of Biological Education (2004) 38(3)

Xylem water transport

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Appendix
Suppliers
LI-COR, inc., 4421 Superior St., Lincoln, N E 6 8 5 0 4 , USA. Tel:
4 0 2 - 4 6 7 - 3 5 7 6 ; Fax: 4 0 2 - 4 6 7 - 2 8 1 9 ; Email: envsales@licor.com.
Jordi Martinez-Vilalta (corresponding author) is currently Research
Fellow at the School of GeoSciences, University of Edinburgh,
Edinburgh EH9 3JU, UK. Tel: +44 131 6505427. Fax: +44 131
6620478. Email: jordi.martinez-vilalta@ed.ac.uk
Josep Pinol is Professor of Ecology at the CREAF/ Ecology Unit at the
Universitat Autbnoma de Barcelona. Email: Josep.Pinol@uab.es.

Journal of Biological Education (2004) 38(3)

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141